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1. Drought suppresses soil predators and promotes root herbivores in mesic, but not in xeric grasslands

   https://doi.org/10.1073/pnas.1900572116  

   Franco, André L.; Gherardi, Laureano A.; de Tomasel, Cecilia M.; Andriuzzi, Walter S.; Ankrom, Katharine E.; Shaw, E. Ashley; Bach, Elizabeth M.; Sala, Osvaldo E.; Wall, Diana H. (June 2019, Proceedings of the National Academy of Sciences)

   Precipitation changes among years and locations along gradients of mean annual precipitation (MAP). The way those changes interact and affect populations of soil organisms from arid to moist environments remains unknown. Temporal and spatial changes in precipitation could lead to shifts in functional composition of soil communities that are involved in key aspects of ecosystem functioning such as ecosystem primary production and carbon cycling. We experimentally reduced and increased growing-season precipitation for 2 y in field plots at arid, semiarid, and mesic grasslands to investigate temporal and spatial precipitation controls on the abundance and community functional composition of soil nematodes, a hyper-abundant and functionally diverse metazoan in terrestrial ecosystems. We found that total nematode abundance decreased with greater growing-season precipitation following increases in the abundance of predaceous nematodes that consumed and limited the abundance of nematodes lower in the trophic structure, including root feeders. The magnitude of these nematode responses to temporal changes in precipitation increased along the spatial gradient of long-term MAP, and significant effects only occurred at the mesic site. Contrary to the temporal pattern, nematode abundance increased with greater long-term MAP along the spatial gradient from arid to mesic grasslands. The projected increase in the frequency of extreme dry years in mesic grasslands will therefore weaken predation pressure belowground and increase populations of root-feeding nematodes, potentially leading to higher levels of plant infestation and plant damage that would exacerbate the negative effect of drought on ecosystem primary production and C cycling. 

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2. Root herbivory controls the effects of water availability on the partitioning between above‐ and below‐ground grass biomass

   https://doi.org/10.1111/1365-2435.13661  

   Franco, André_L_C; Gherardi, Laureano_A; de_Tomasel, Cecilia_M; Andriuzzi, Walter_S; Ankrom, Katharine_E; Bach, Elizabeth_M; Guan, Pingting; Sala, Osvaldo_E; Wall, Diana_H; Holeski, ed., Liza (August 2020, Functional Ecology)

   Abstract The fraction of primary productivity allocated below‐ground accounts for a larger flow of carbon than above‐ground productivity in most grassland ecosystems. Here, we addressed the question of how root herbivory affects below‐ground allocation of a dominant shortgrass prairie grass in response to water availability. We predicted that high levels of root herbivory by nematodes, as seen under extreme drought in sub‐humid grasslands, would prevent the high allocation to root biomass normally expected in response to low water availability.We exposed blue gramaBouteloua gracilis, which accounts for most of the net primary productivity in the shortgrass steppe of the central and southern Great Plains, to three levels of water availability from extreme low to intermediate and extreme high crossed with a gradient of root­herbivore per cent abundance relative to the total nematode community in soil microcosms.As hypothesized, the effect of water availability on below‐ground biomass allocation was contingent on the proportion of root herbivores in the nematode community. The relationship between below‐ground biomass allocation and water availability was negative in the absence of root herbivory, but tended to reverse with increasing abundance of root feeders. Increasing abundance of root‐feeding nematodes prevented grasses from adjusting their allocation patterns towards root mass that would, in turn, increase water uptake under dry conditions. Therefore, below‐ground trophic interactions weakened plant responses and increased the negative effects of drought on plants.Our work suggests that plant responses to changes in precipitation result from complex interactions between the direct effect of precipitation and indirect effects through changes in the below‐ground trophic web. Such complex responses challenge current predictions of increasing plant biomass allocation below‐ground in water‐stressed grasslands, and deserve further investigation across ecosystems and in field conditions. A freePlain Language Summarycan be found within the Supporting Information of this article. 

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3. Nutrient addition increases grassland sensitivity to droughts

   https://doi.org/10.1002/ecy.2981  

   Bharath, Siddharth; Borer, Elizabeth T.; Biederman, Lori A.; Blumenthal, Dana M.; Fay, Philip A.; Gherardi, Laureano A.; Knops, Johannes M. H.; Leakey, Andrew D. B.; Yahdjian, Laura; Seabloom, Eric W. (February 2020, Ecology)

   Abstract Grasslands worldwide are expected to experience an increase in extreme events such as drought, along with simultaneous increases in mineral nutrient inputs as a result of human industrial activities. These changes are likely to interact because elevated nutrient inputs may alter plant diversity and increase the sensitivity to droughts. Dividing a system’s sensitivity to drought into resistance to change during the drought and rate of recovery after the drought generates insights into different dimensions of the system’s resilience in the face of drought. Here, we examine the effects of experimental nutrient fertilization and the resulting diversity loss on the resistance to and recovery from severe regional droughts. We do this at 13 North American sites spanning gradients of aridity, five annual grasslands in California, and eight perennial grasslands in the Great Plains. We measured rate of resistance as the change in annual aboveground biomass (ANPP) per unit change in growing season precipitation as conditions declined from normal to drought. We measured recovery as the change in ANPP during the postdrought period and the return to normal precipitation. Resistance and recovery did not vary across the 400‐mm range of mean growing season precipitation spanned by our sites in the Great Plains. However, chronic nutrient fertilization in the Great Plains reduced drought resistance and increased drought recovery. In the California annual grasslands, arid sites had a greater recovery postdrought than mesic sites, and nutrient addition had no consistent effects on resistance or recovery. Across all study sites, we found that predrought species richness in natural grasslands was not consistently associated with rates of resistance to or recovery from the drought, in contrast to earlier findings from experimentally assembled grassland communities. Taken together, these results suggest that human‐induced eutrophication may destabilize grassland primary production, but the effects of this may vary across regions and flora, especially between perennial and annual‐dominated grasslands. 

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4. Direct and indirect effects of nitrogen enrichment on soil organisms and carbon and nitrogen mineralization in a semi‐arid grassland

   https://doi.org/10.1111/1365-2435.132  

   Chen, Dima; Xing, Wen; Lan, Zhichun; Saleem, Muhammad; Wu, Yunqiqige; Hu, Shuijin; Bai, Yongfei (January 2019, Functional ecology)

   Semi‐arid grasslands on the Mongolian Plateau are expected to experience high inputs of anthropogenic reactive nitrogen in this century. It remains unclear, however, how soil organisms and nutrient cycling are directly affected by N enrichment (i.e., without mediation by plant input to soil) vs. indirectly affected via changes in plant‐related inputs to soils resulting from N enrichment. To test the direct and indirect effects of N enrichment on soil organisms (bacteria, fungi and nematodes) and their associated C and N mineralization, in 2010, we designated two subplots (with plants and without plants) in every plot of a six‐level N‐enrichment experiment established in 1999 in a semi‐arid grassland. In 2014, 4 years after subplots with and without plant were established, N enrichment had substantially altered the soil bacterial, fungal and nematode community structures due to declines in biomass or abundance whether plants had been removed or not. N enrichment also reduced the diversity of these groups (except for fungi) and the soil C mineralization rate and induced a hump‐shaped response of soil N mineralization. As expected, plant removal decreased the biomass or abundance of soil organisms and C and N mineralization rates due to declines in soil substrates or food resources. Analyses of plant‐removal‐induced changes (ratios of without‐ to with‐plant subplots) showed that micro‐organisms and C and N mineralization rates were not enhanced as N enrichment increased but that nematodes were enhanced as N enrichment increased, indicating that the effects of plant removal on soil organisms and mineralization depended on trophic level and nutrient status. Surprisingly, there was no statistical interaction between N enrichment and plant removal for most variables, indicating that plant‐related inputs did not qualitatively change the effects of N enrichment on soil organisms or mineralization. Structural equation modelling confirmed that changes in soil communities and mineralization rates were more affected by the direct effects of N enrichment (via soil acidification and increased N availability) than by plant‐related indirect effects. Our results provide insight into how future changes in N deposition and vegetation may modify below‐ground communities and processes in grassland ecosystems. 

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5. Trophic group specific responses of alpine nematode communities to 18 years of N addition and codominant plant removal

   https://doi.org/10.1007/s11104-023-06281-3  

   Shepherd, Rachel M.; Brigham, Laurel M.; de Mesquita, Clifton P.; Gattoni, Kaitlin M.; Gendron, Eli M.; Hahn, Philip G.; Schmidt, Steven K.; Smith, Jane G.; Suding, Katharine N.; Porazinska, Dorota L. (October 2023, Plant and Soil)

   Background and Aims Human-driven nitrogen (N) deposition can alter soil biogeochemistry and plant communities, both critical to soil biota. However, understanding the relative impact of the relationship between nutrient resources and plants on soil communities has been hindered by a lack of experimental manipulations of both factors. We hypothesized that soil nematode communities would be structured predominantly by N addition via overall increased abundance, decreased diversity, and compositional shifts to dominance of r-selected bacterial-feeding nematodes. In contrast, we expected plant efects to be less evident and restricted to nematodes directly associated with plants. Methods We used a long-term (18-yrs) experiment in moist meadow alpine tundra involving N addition and codominant plant (nitrophilic Deschampsia cespitosa and nitrogen sensitive Geum rossii) removal. We characterized nematode communities via 18S rRNA metabarcoding and used soil biogeochemistry, plant, and microbial variables to determine factors shaping their communities. Results The N addition treatment increased overall nematode abundance, decreased diversity, and afected the composition of all nematode trophic groups. Overall, nematode communities shifted to dominance of bacterial feeding nematode taxa adapted to N-enriched environments. The likely drivers of this shift were increased soil nitrate and lower pH. The direct efects of codominant plants were more limited, with only changes in Geum rossii appearing to afect nematode responses. Conclusion Overall, nematode communities in N-limited alpine ecosystems are highly sensitive to increases in N availability, irrespective of the nature of N preferences of codominant plants. The resulting nematode community restructuring could signify future shifts in soil functioning throughout alpine landscapes. 

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